molecules
Review
Quality Chemistry, Physiological Functions, and Health
Benefits of Organic Acids from Tea (Camellia sinensis)
Hongbo Chen 1,2,† , Fei Yu 2,† , Jiaxin Kang 2 , Qiao Li 1 , Hasitha Kalhari Warusawitharana 2, * and Bo Li 2, *
Citation: Chen, H.; Yu, F.; Kang, J.; Li,
Q.; Warusawitharana, H.K.; Li, B.
Quality Chemistry, Physiological
Functions, and Health Benefits of
Organic Acids from Tea (Camellia
sinensis). Molecules 2023, 28, 2339.
https://doi.org/10.3390/
molecules28052339
Academic Editor: Jason Tze
Cheng Tzen
Received: 7 February 2023
Revised: 25 February 2023
Accepted: 27 February 2023
Published: 3 March 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
This article is an open access article
distributed under the terms and
conditions of the Creative Commons
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Molecules 2023, 28, 2339. https://doi.org/10.3390/molecules28052339
1 Modern Service Industry Research Institute, Zhejiang Shuren University, Hangzhou 310015, China
2 Department of Tea Science, Zhejiang University, Hangzhou 310058, China
* Correspondence: 11816127@zju.edu.cn (H.K.W.); drlib@zju.edu.cn (B.L.)
† These authors contributed equally to this work.
Abstract: Organic acids account for around 3% of the dry matter in tea leaves, and their composition
and contents vary in different types of tea. They participate in the metabolism of tea plants, regulate
nutrient absorption and growth, and contribute to the aroma and taste quality of tea. Compared
with other secondary metabolites in tea, the researches on organic acids are still limited. This
article reviewed the research progresses of organic acids in tea, including analysis methods, the
root secretion and physiological function, the composition of organic acids in tea leaves and related
influencing factors, the contribution of organic acids to sensory quality, and the health benefits, such
as antioxidation, promotion of digestion and absorption, acceleration of gastrointestinal transit, and
regulation of intestinal flora. It is hoped to provide references for related research on organic acids
from tea.
Keywords: tea; organic acids; physiological function; sensory quality; health benefit
1. Introduction
Organic acids are a class of organic compounds with acidity that originates from the
carboxyl group (-COOH), sulfonic group (-SO3 H), sulfinic group (-RSOOH), and carboxylic
group (-RCOSH), etc. Carboxylic acids are the most common type of organic acids. They
are divided into aliphatic, aromatic, and terpenoid compounds based on their structures
and into monocarboxylic acids, dicarboxylic acids, and polycarboxylic acids according
to the number of carboxylic groups [1]. Organic acids are the intermediate products of
carbohydrates in biochemical reactions and are the participants in plant metabolic activities.
They are also important intermediates in metabolic pathways, such as the tricarboxylic acid
cycle and shikimic acid pathway, and play a vital role in energy transfer and maintenance
of cell osmotic pressure [2,3]. Organic acids in tea are mainly aliphatic acids (such as oxalic
acid, malic acid, and citric acid) and aromatic acids (such as benzoic acid, salicylic acid, and
cinnamic acid). Until now, more than 40 organic acids have been discovered in different tea,
including more than 10 in tea soup and over 30 in its aroma. Organic acids were thought to
contribute to the tea flavor, epically the sourness of tea soup, and have a variety of health
benefits for the human body, such as regulation of intestinal flora and antioxidation. In
addition, organic acids influence nutrient absorption of tea plants, such as phosphorus and
aluminum ions. Compared with other metabolites of tea plants, such as catechin, theine,
theanine, and flavonoid glycosides, there are few studies on organic acids in tea [4]. In this
paper, the detection method of organic acids, their composition and function in tea roots
and leaves, the influence of the plant variety and processing technology on them, as well as
their health functions, are reviewed to provide references for future studies.
2. Detection Method of Organic Acids
The titration method, atomic absorption method, gas chromatography, ion exchange
chromatography, high-performance capillary electrophoresis, and high-performance liquid
https://www.mdpi.com/journal/molecules
Molecules 2023, 28, 2339 2 of 11

chromatography are the main methods to detect the organic acids in tea. He et al. [5]
introduced the principle, application, advantages, and disadvantages of the above detection
methods, according to which this paper makes a further summary, as shown in Table 1.
The most commonly used method for the analysis of organic acids is high-performance
liquid chromatography (HPLC), which mostly uses the C18 column as the chromatographic
column and tends to add phosphoric acid to the mobile phase. Phosphoric acid is more
acidic than formic acid and acetic acid, which are commonly used in high-performance
liquid chromatography mobile phase. Plus, it has no UV background or volatility with
moderate ionic strength. However, owing to the generally small molecular weight and
high polarity of organic acids, the separation degree of organic acids on HPLC is usually
low. In addition, the phosphate in the mobile phase is prone to crystallize and tends to
block the instrument pipeline, causing damage to instruments and thus increasing the cost
of equipment operation and maintenance [6–10].

Table 1. Analysis methods of organic acids.

Methods Principle Characteristics

Titration end points are determined by
Titration
indicators or potential changes.
Atomic-absorption
Oxalate is precipitated into calcium oxalate.
spectrophotometry
The samples are separated according to the
GC-MS difference in boiling point, polarity,
and adsorbability.
Separation is achieved according to the
Ion exchange difference in the ability of the separated
chromatography components to undergo ion exchange with
the stationary phase.
High-performance capillary Separation is based on the electrophoretic
electrophoresis differences between components in a sample.
Each component has different partition
HPLC-UV
coefficients in the two phases.
Determination of total, not individual organic
acid. Convenient with low sensitivity.
Determination of oxalic acid.
Qualitative and quantitative analysis of small
organic acid molecules, but not applicable for
large molecules.
Not suitable for samples with
complex composition.
High-separation efficiency. Less sample
loading and relatively poor repeatability.
High-separation efficiency. Less sample
loading and the use of phosphates, which is
easy to form crystals to block the pipeline.

With the development of modern analytical technology, more and more researchers
use ultra-high liquid chromatography-mass spectrometry (UPLC-MS) technology to detect
organic acids and quantitative analysis carried out under multi-reaction detection mode
(MRM) [11–18]. The mobile phase of this method does not need to use phosphoric acid,
but mainly uses formic acid or acetic acid, water and acetonitrile. The method also enjoys
exquisite specificity, avoiding the qualitative and quantitative interference caused by the
close peak time and low separation degree between different organic acids. Plus, the
sensitivity and accuracy of the method are also improved. Nowadays, this technique has
been used for the detection of organic acids in fruits, tobacco leaves, and Chinese medicinal
herbs. The author summarizes the MRM detection conditions of common organic acids in
tea, such as fragment ion pair, cone hole voltage, and collision pressure [17–19] (Table 2),
all using electrospray ionization (ESI) and negative ion mode. In this way, the article hopes
to provide a reference for the establishment of the method for the detection of organic acids
in tea leaves.
 Organic
medicinal
Table
organic
detection ofherbs.
2. Determination
acids in2), The
tea,
organic author
conditions
such summarizes
ofleaves.
major
aselectrospray
fragment ionPrecursor
the
organic MRM
acids
pair, cone inDaughter
detection
tea
hole under Ion Cone
conditions
MRM-MS.
voltage, and of common
collision pressure
[17–19]
way, the (Table
article allacids
hopes using
to in tea
provide
Formula ionization
a reference for the (ESI) and
establishmentnegative
of ion
the mode.
method In (V)
for
Collision this
the
Organic
medicinal
Acids
Table
organic2. herbs.
Determination
acids in The
tea, author
conditions
such as summarizes
of
fragmentmajor Precursor
Ionsthe
(m·z
organic
ion MRM
−1
acids
pair, cone) Daughter
in detection
tea
hole(m·z −1
under ) Ion
voltage, Cone
conditions
(V)
MRM-MS.
and of
collision common
pressure
[17–19]
way, the(Table
detection article 2), allacids
hopes
of organic usinginelectrospray
Formula
to provide ionization
a reference
tea leaves. for the(ESI) and negative
establishment ionmethod
of the mode. for
CollisionIn (V)
this
the
Organic
Acids Precursor
Ions (m·z −1) Daughter −1)Ion Cone
organic
[17–19]
Table
way, the
detection
acids
2. (Table in2),hopes
Determination
article
of organic
tea, such
allacids
using aselectrospray
Formula fragment
conditions
in teaofleaves.
to provide majorion pair,
ionization
organic
a reference cone
acids
for the tea(m·z
hole
(ESI) voltage,
and
inestablishment
under and
negative (V)ion
MRM-MS.
of thecollision
mode.pressure
Collision
method In this
for(V)
the
Organic
Acids
[17–19] (Table 2), all using electrospray Precursor
Ions (m·z
ionization
−1) Daughter
(ESI) (m·z
and
−1) Ion Cone
negative(V) ion mode. In this
way, the
detection
Table article
of hopes
organic
2. Determination to provide
Formula
acids in tea
conditions a reference
ofleaves.
major organicfor the
acids in tea 85.0
establishment
under * MRM-MS.
of the method for
Collision the
(V)
Quinic
way, Acids
Organic
the acid
article hopes to provide amajor Ions 191
(m·z
Precursor
reference for
−1)
the (m·z−1*) Ion of
Daughter 35
(V)
Cone 20for the
detection
Table
Quinic
Molecules
of organic
2. Determination
2023, acid
28, 2339
acids in teaof
Formula
conditions leaves. organic
191 acids inestablishment
tea85.093 MRM-MS.
under the method
35 Collision
20 (V) 3 of 11
Acids
Organic
detection
Table 2. of organic acids
Determination in teaof
conditions leaves.
major Ions
organic (m·z
Precursoracids
−1) in (m·z
Daughter
tea85.0
93
under* )Ion
−1 (V)
Cone
MRM-MS.
Quinic acid Formula 191 −1 35 Collision
20 (V)
TableOrganic
Acids
2. Determination conditions Precursor
Ions
of major organic (m·zacids) in Daughter
tea(m·z93 −1*) Ion
85.0
under Cone
(V)
MRM-MS.
Quinic
Organic acid Formula 191
Precursor Daughter 35 Collision
20 (V)
Table Acids
2. Determination conditions of major organic
Formula Ions (m·z
acids−1) in (m·z93 −1**) Ion
tea114.9
under
85.0 Cone
MRM-MS.(V) Collision (V)
Quinic
Organic
Malic
Acids acid
acid Ions
Table 2. Determination191
Precursor
133
(m·z −1) Daughter
conditions (m·z
of −1)Ionorganic35
Cone
49 acids in tea
(V) 20 under MRM-MS.
16
Formula 93major
114.9
85.0 ** Ion Cone Collision
Organic
Malic
Quinic
Acids acid
acid Precursor
133
Ions191
(m·z−1) Daughter
(m·z
45
114.9
−1) 49
35
(V) 16 (V)
20
Formula 85.0
45
93 * Collision (V)
Malic
Acids
Quinic
Organic acid
acid
Acids Formula Ions133(m·z
191
Precursor Ions) (m·z114.9
−1 (m·z
−1 ) ) Daughter
−1 49
35 Ion (m·z−16
(V) 20
1) Cone (V) Collision (V)
85.0
45
93 *
*
Malic acid
Quinic acid 133
191 49
35 16
20
85.0
93 **
45
110.0
114.9
Quinic
Citric
Malic acidacid 191
191.2
133 85.0 * 35
50
49 20
18
16
Quinic acid 191191 110.0
45 **
93
87
114.9 3585.0 *
20
Quinicacid
Citric acid 191.2 50 18 35 20
Malic acid 133 93 **
110.0
87 49 93 16
Citric 114.9
45
Malic acid
acid 191.2
133 110.0
87
114.9 **
50
49 18
16
Citric acid
Malic acid 191.2
133 45 50
49 18
16
87
110.0
73.1
45 **
114.9 114.9 *
Citric
Succinic
Malic acid
Malic acidacid 191.2
116.9
133133 114.9
73.1 ** 50
60
49 18
19
16 49 16
Malic acid
Succinic acid
acid 133
116.9 87 *
99.2
110.0
45 4945
60 16
19
Citric 191.2 73.1
45 *
99.2 * 50 18
Succinic acid 116.9 110.0
87 60 19
Citric acid 191.2 73.1
99.2
110.0 ** 50 18
Succinic
Citric acid
acid
Citric acid 116.9
191.2 191.2
87 110.0 *
60
50 19
18 50 18
99.2
73.1
87 ***
110.0
70.9 87
Succinic
Citric acid
Fumaric acid
Acid 116.9
191.2
114.9 110.0 * 60
50
68 19
18
11.5
Citric acid 191.2 99.2
87 *
73.1
70.9
97.8 50 18
Succinic
Fumaric acid
Acid 116.9
114.9 87 **
70.9 60
68 19
11.5
73.1
99.2
97.8 73.1 *
Fumaric
Succinic Acid
Succinic acid
acid 114.9
116.9 116.9 68
60 11.5
19 60 19
73.1
70.9
99.2*
97.8 99.2
Succinic Acid
Fumaric acid 116.9
114.9 73.1 60
68 19
11.5
Succinic Acidacid 116.9 99.2**
97.8
70.9 60 19
Fumaric 114.9 73.1
102.9 * 68 11.5
Succinic
Tartaric acid
acid 116.9
149 97.8**
99.2
70.9 60
56.970.9 * 19
15.3
Fumaric Acid
Fumaric Acid 114.9 114.9 99.2
102.9
73 * 68 11.5 68 11.5
Tartaric acid 149 70.9
97.8 ** 56.9 97.8 15.3
Fumaric Acid 114.9 102.9
73 68 11.5
Tartaric acid 149 70.9
97.8* 56.9 15.3
Fumaric Acid 114.9 102.9
73 **
70.9 68 11.5
Tartaric
Fumaric Acid acid 149
114.9 97.8 56.9
68 15.3
11.5
73
102.9
97.8**
70.9 102.9 *
Tartaric
Fumaric acid
Acid
Tartaric acid 149149
114.9 43.2 ** 56.9
68 15.3
11.5 56.9 15.3
73
97.8
102.9 73
Lactic acid
Tartaric acid 89
149 43.2 * 63
56.9 21
15.3
Lactic acid 89 70.9
102.9
73 * 63 21
Tartaric acid 149 43.2
70.9
102.9 * * 56.9 15.3
Lactic acid
Tartaric acid 89
149 73 63
56.9 21
15.3
43.2
70.9
102.9 * 43.2 *
Lactic
Tartaric acid
Lactic acid
acid 89 89
149 73 * 63
56.9 21
15.3 63 21
70.9
102.9
43.2 ** 70.9
Tartaric
Lactic acidacid 149
89 73 56.9
63 15.3
21
73 *
70.9
43.2
Gallic acid
Lactic acid 168.9
89 78.8 * 77
63 29
21
Gallic acid 168.9 43.2
70.9
78.8 *
* 77 29
Lactic acid 89 43.2 * 63 21
Gallic
Gallic acid
acid
Lactic acid 168.9
89 168.9 70.9
78.8 * 7778.8
63 * 29
21 77 29
Gallic acid
acid 168.9 43.2
70.9**
78.8 77 29
Lactic 89 43.2 63 21
Lactic
Gallic acid 89
168.9 70.9**
78.8 63
77 21
29
43.1
70.9*
Pyruvic
Gallic acidacid 87
168.9 78.8 50 12.1
OR PEER REVIEW
Pyruvic
Pyruvic acidacid 87 87 68.7*
43.1 77
5043.1 * 294 of 12
12.1
OR PEER REVIEW Gallic acid 168.9 43.1
68.7**
78.8 7768.7 294 of 12 50 12.1
Pyruvic acid 87 50 12.1
Gallic acid
x FOR PEER REVIEW 168.9 78.8
43.1
68.7 * 77 29 4 of 12
Pyruvic
OR PEER REVIEW Gallic acidacid 87
168.9 78.8 50 12.1
294 of 12
68.7**
43.1 77
Pyruvic
Gallic acidacid 87
168.9 78.8
87 * 5087
77 12.1
29
Ascorbic acid
Ascorbic acid 175175 68.7
43.1
87 ** 56 19.1 56 19.1
Pyruvic acid 87 114.9 114.9 *
50 12.1
Ascorbic acid 175 93.0
43.1
68.7 * 56 19.1
Shikimic
Pyruvic acid acid 173
87 87 *
114.9 65.4
50 19.4
12.1
Ascorbic acid 175 43.1
136.8
93.0
68.7 * 56 19.1
Pyruvic
Shikimic acidacid
acid 87
173 87
114.9 * 50
65.4 12.1
19.4
Ascorbic 175 43.1 * *
93.0
68.7
136.8 56 19.1
Pyruvic
Shikimic acidacid 87173 114.9
87
43.1
93.0 ** 50 65.4 12.1 19.4
Ascorbic acid 175 68.7
136.8 5693.0 * 19.1
Pyruvic
Shikimic acid
Shikimic acid
acid 87 173
173 114.9
87 * 50
65.4 12.1
19.4 65.4 19.4
68.7
136.8 136.8
Ascorbic acid 175 56 19.1
87 *
114.9
Ascorbic
Chlorogenic acid 175 87 ** 56 19.1
Ascorbic acid 353.1
175 114.9
191.1 17
56 22
19.1
acid
Chlorogenic 87 *
114.9
Ascorbic acid
Chlorogenic 175
353.1 191.1
87 * * 56
17 19.1
22
acid acid
Ascorbic
Chlorogenic acid 175 353.1
353.1
114.9191.1 * 56 17
191.1 * 19.1 22 17 22
Chlorogenic 114.9
acid 353.1 191.1 ** 17 22
acid
Cinnamic
146.9 103.0 * 48 14
acid
Cinnamic
Cinnamic
Cinnamicacid 146.9 146.9 103.0 * 48
103.0 * 14 48 14
acid 146.9 103.0 * 48 14
Cinnamic
acid 146.9 103.0 * 48 14
acid
Benzoic acid
Benzoic acid 120.9
120.9 77.0 * 40
77.0 * 17 40 17
Benzoic acid 120.9 77.0 * 40 17
Benzoic acid 120.9 77.0 * 40 17
Benzoic acid 120.9 77.0 * 40 17

Salicylic acid 137 92.9 * 48 21

Salicylic acid 137 92.9 * 48 21
Salicylic acid 137 92.9 * 48 21
*Salicylic
Represents quantitative ion.
acid 137 92.9 * 48 21
 Cinnamic
146.9 103.0 * 48 14
acid
Molecules 2023, 28, 2339 4 of 11

Benzoic acid Table 2. Cont. 120.9 77.0 * 40 17

Organic Acids Formula Precursor Ions (m·z−1 ) Daughter Ion (m·z−1 ) Cone (V) Collision (V)

Salicylic acid
Salicylic acid 137137 92.9 * 48
92.9 * 21 48 21

* Represents quantitative ion.* Represents quantitative ion.

3. Secretion and Function of Organic Acids in Tea Roots
3. Secretion and Function of Organic Acids in Tea Roots
The types and contents of organic acids secreted by the root system of tea plants
The types and contents of organic acids secreted by the root system of tea plants are
are significantly different. The influencing factors mainly include varieties, seasons, light
significantly different. The influencing factors mainly include varieties, seasons, light
conditions, soil pH, etc. The organic acids secreted by roots are of great significance to the
conditions, soil pH, etc. The organic acids secreted by roots are of great significance to the
absorption of nutrients and the normal growth of tea plants.
absorption of nutrients and the normal growth of tea plants.
3.1. Types and Contents of Organic Acids in Tea Roots
3.1. Types and Contents of Organic Acids in Tea Roots
The organic acids secreted by the root system of the tea plant mainly include tartaric
The organic acids secreted
acid, by the acid,
oxalic root system of the
malic acid, teaacid,
citric plantandmainly include
succinic acid tartaric
[20–22]. The types and contents of
acid, oxalic acid, malic acid, citric acid,
organic acidsand succinic
secreted acid [20–22].
by roots vary in The typesseasons
different and contents
and varieties. For the tea variety
of organic acids secreted by roots vary
Longjing 43, it in
was different
found thatseasons and varieties.
in spring, citric acidFor the tea the most of all the organic
is secreted
variety Longjing 43, it wasacids
found that µg/individual
(1007 in spring, citricplant),
acid isandsecreted
oxalicthe
acidmost
and of all the
malic acid are the least. In summer,
organic acids (1007 μg/individual
succinicplant),
acid isand oxalicthe
secreted acid
most and(1500
malic acid are the least.
µg/individual plant),In and oxalic acid is the least.
summer, succinic acid is secreted the most (1500 μg/individual plant), and oxalic
In autumn, the secretion of oxalic acid is the highest (388 µg/individual acid is plant), and the
the least. In autumn, the secretion
secretionofofoxalic acidisisthe
citric acid thelowest.
highestIn(388termsμg/individual plant), of oxalic acid, succinic acid,
of the total amount
and the secretion of citric acid
malicisacid,
the lowest.
and citric Inacid,
terms theofsecretion
the totalisamount of oxalic
the highest in summeracid, (1856 µg/individual plant)
succinic acid, malic acid, and least
citric inacid, the secretion
autumn is the highest
(444 µg/individual in summer
plant). The three (1856
different tea varieties, Biyun,
Fuding,
μg/individual plant) and least and Longjing
in autumn 43, have different
(444 μg/individual plant).total
Theorganic acid secretion
three different tea in autumn, with Fuding
varieties, Biyun, Fuding, andthe Longjing
most (1213 have differentplant)
43,µg/individual total and Biyun
organic thesecretion
acid least (50in µg/individual plant). Light
autumn, with Fuding the most (1213 μg/individual plant) and Biyun the least (50 product of metabolism,
is essential for the secretion of organic acids. As an intermediate
μg/individual plant). Light organic acids arefor
is essential mostly
the produced
secretion inofthe processacids.
organic of the Astricarboxylic
an acid cycle, and their
secretion isorganic
intermediate product of metabolism, inseparable acidsfrom the photosynthesis
are mostly produced inofthe plants.
process Under
of light conditions, the total
the tricarboxylic acid cycle,amount
and their of secretion
organic acids secreted by
is inseparable thethe
from roots of Longjing 43
photosynthesis of and the Fuding tea tree is
muchthe
plants. Under light conditions, higher
total than
amount thatofunder dark
organic conditions,
acids secretedwhile
by the oxalic
rootsacidof is scarcely secreted under
Longjing 43 and the Fudingdark conditions
tea tree is much[21].higher than that under dark conditions, while
oxalic acid is scarcely secreted under et
Pang darkal. conditions
found that [21].
tea polyphenols could induce the secretion of malic acid and
citric acid from the root
Pang et al. found that tea polyphenols could induce the system of the tea plant.
secretion Additionally,
of malic acid and a proper amount of water-
soluble
citric acid from the root system material
of the trimmings
tea plant. can induce
Additionally, the secretion
a proper amount of of oxalic
water-acid, malic acid, citric acid,
soluble material trimmingsand cansuccinic
induce acid, thus reducing
the secretion the pH
of oxalic acid,value
malicofacid,
the collected solution. Therefore, the water-
citric acid,
soluble components of tea plant trimming may be one of the causes of soil acidification
in tea plantations. Soil pH has a major effect on the secretion of organic acids from tea
plants [22]. Tang et al. took Longjing 43 as the material and found that the concentration of
oxalic acid in roots and that of malic acid, citric acid, and oxalic acid in mature leaves are
increased at pH 6.0 relative to the condition of pH 4.0 and pH 5.0 [23].

3.2. Organic Acids in Tea Roots and Nutrient Absorption

The organic acids secreted by the root system of tea tree can improve tea plants’
absorption of phosphorus elements, reduce the adsorption of fluorine by the iron film
on the root surface of tea plants and play a significant role in the healthy growth of tea
plants in a high-aluminum acid environment. The organic acids secreted by tea plants
during their growth process can dissolve insoluble inorganic phosphorus in the soil. After
organic acid treatment, the content of insoluble inorganic phosphorus in soil decreases
to varying degrees. Among them, oxalic acid has the strongest effect, followed by citric
acid and malic acid, and succinic acid has the weakest solubility [21]. The secretion of
organic acids from roots may be one of the main reasons for the strong ability of tea plants
to absorb phosphorus. Tea is a typical plant with high enrichment of fluorine elements,
Molecules 2023, 28, 2339 5 of 11

especially in crude and old tea leaves. The fluorine element is mainly ingested from the
roots of the tea plant and accumulated in the leaves [24,25]. The content of Fe2+ and Al3+
in the soil of a tea garden is high, and thus the iron oxide film is easy to form on the root
surface of a tea tree, which will promote the absorption of fluorine by a tea tree. With the
increase of organic acid concentration, the adsorption capacity of root surface iron film on
fluorine decreases [26]. Liu et al. conducted hydroponic experiments on tea seed seedlings
of the Fuding white tea variety and found that under the stimulation of aluminum ions,
the roots mainly secreted oxalic acid, malic acid, and citric acid, accounting for 85–93% of
the total amount of organic acid secreted. The secretion of oxalic acid would increase with
the concentration of aluminum ions rose. Therefore, oxalic acid may be able to balance the
concentration of aluminum ions in tea tree roots. Given the fact that tea trees can still grow
well in a high-aluminum acidic environment, it is likely that organic acids play an active
role in alleviating aluminum toxicity [27,28].
Organic acids in the root system of tea trees are vital to the material circulation in the
soil of tea gardens. Lin et al. chose Fujian Jiukeng seedlings as experimental materials
and found that under the environment of Pb2+ and Cd2+ , the composition of organic acids
secreted by roots did not change, mainly including oxalic acid, malic acid, lactic acid, acetic
acid, citric acid, and succinic acid. However, the amount of each acid changes to varying
degrees. Among them, the secretion of succinic acid and malic acid increases significantly,
while citric acid decreases noticeably. This may be an important mechanism for the complex
detoxification of organic acids and metal ions [29]. Zeng et al. found that organic acids
such as citric acid, malic acid, tartaric acid, and oxalic acid can activate F, P, Zn, Fe, Al, Cu,
and Mn elements in the soil [30].

4. Organic Acids in Tea

The composition and content of organic acids in tea leaves are closely related to the
type, processing technology, variety, and growth period of tea [7,31–36].
Tukhvatshin et al. showed that the organic acids in Fu’an white tea leaves are mainly
citric acid, gallic acid, benzoic acid, oxalic acid, salicylic acid, ascorbic acid, and chlorogenic
acid. Among them, the content of citric acid was the highest, p-coumaric acid and fumaric
acid were the lowest, and succinic acid was not detected. The total amount of organic
acids in white tea processing ranged from 5.6839 to 10.1014 mg/g, and the order is crude
tea > 24H > 48H > 32H > 40H > 16H > 8H > fresh leaf. The contents of citric acid, gallic
acid, oxalic acid, caffeic acid, cinnamic acid, and acetic acid show an increasing trend.
Among them, the contents of citric acid, gallic acid, and oxalic acid in crude white tea are
4.7873, 2.4122, and 1.2758 mg/g, which are respectively 2.9, 1.4, and 1.7 times those in fresh
leaves. During the processing of white tea, the contents of chlorogenic acid and p-coumaric
acid show a decreasing trend. The contents of chlorogenic acid and p-coumaric acid in fresh
leaves are 0.1614 and 0.0366 mg/g, 1.6 and 3.0 times those of white tea, respectively. There
is no obvious change in the content of benzoic acid, salicylic acid, ferulic acid, fumaric acid,
and ascorbic acid content. The content of benzoic acid peaked eight hours after withering,
ascorbic acid 16 h, salicylic acid 24 h, and ferulic acid 48 h [31]. He et al. conducted an
analysis of the components of organic acids in Silver Needle Pekoe, White Peony, and
Shou-mee and found that with the decrease of leaf freshness, the number of organic acids
in white tea decreased, among which the content of lactic acid, acetic acid, citric acid, and
fumaric acid gradually decreases. However, the content of malic acid increases with the loss
of freshness, while oxalic acid and ascorbic acid have no distinct change pattern. Overall,
acetic acid is the main organic acid in the three kinds of white tea samples, accounting for
32.8%, 46.5%, and 42.5% of the total organic acid [6].
During black tea processing, the total amount of organic acids, including oxalic acid,
citric acid, succinic acid, malic acid, etc., significantly increases after withering, rolling, and
fermentation. The contents of organic acids are 1.44%, 1.51%, and 1.72% in fresh leaves,
withered leaves, and rolled leaves, respectively. After drying, the organic acid content in
the tea leaves will decrease [35]. Yu et al. explored dynamic changes of organic acids during
Molecules 2023, 28, 2339 6 of 11

black tea processing by a UPLC-MRM-MS method. The results showed that the contents
of quinic acid and citric acid increased and then decreased during the withering process,
but there was no obvious change during the whole process. Malic acid content obviously
decreased at the early stage of withering, and gallic acid content markedly increased after
rolling [37]. Xie et al. found that the total amount of organic acids in Keemun black
tea stored at room temperature for 6 years considerably increased, but the tendency of
each organic acid compound was different. For example, the content of oxalic acid and
acetic acid greatly increased, while the citric acid significantly decreased, leading to the
overall sour and poor taste of tea soup [33]. Mao analyzed 32 Gongfu black teas produced
in 14 provinces and used HPLC to determine eight organic acids, including oxalic acid,
pyruvic acid, L-malic acid, L-ascorbic acid, lactic acid, acetic acid, citric acid, and succinic
acid. It was found that the content of L-malic acid in Gongfu black tea samples was the
highest, followed by oxalic acid and succinic acid, and the content of acetic acid was the
lowest, which was only detected in some samples. The content of L-malic acid and acetic
acid in different types of Gongfu black teas greatly varies [38]. Zhang et al. analyzed six
black teas from different origins, including Ceylon black tea from Sri Lanka, Ninghong
from Jiangxi, Dianhong from Yunnan, Yinghong from Guangdong, Keemun black tea from
Anhui and Shimen black tea from Hunan. A total of 11 organic acids are detected: oxalic
acid, tartaric acid, formic acid, pyruvic acid, malic acid, ascorbic acid, lactic acid, acetic
acid, citric acid, succinic acid, and fumaric acid, among which oxalic acid and acetic acid
are the main organic acids [10].
Li et al. detected nine organic acids, namely oxalic acid, tartaric acid, pyruvate, malic
acid, acetic acid, citric acid, succinic acid, fumaric acid, and α-ketoglutaric acid, during the
flowering process of dark tea. It is found that the amount of malic acid decreases by 85.8%,
while the content of succinic acid increases by 8.42 times that of before fermentation [39].
Ding found that the total amount of organic acids in Fuzhuan brick tea processing increased
from 42.63 mg/g in raw tea to 43.44 mg/g in the finished tea. After 5 days of flowering, the
content of D-malic acid and α-ketoglutarate acid decreases, while ascorbic acid, acetic acid,
and citric acid increase [40].
Liu studied the content of organic acids in different leaf positions of one-bud four-leaf
Cuifeng species, Longjing 43, Jiukeng, Huangjingui, Yunkang 10, and Fuding 5, as well
as representative teas on sale, such as black tea, green tea, oolong tea, and Pu ‘er tea. He
also studied the leaching characteristics of organic acids in Wuyi green tea under different
brewing conditions. In terms of the leaf position, the total amount of organic acid decreases
with the decrease in fresh leaf tenderness. The content of oxalic acid decreases significantly,
and the content of organic acids in the fourth leaf is only 49.5% of that in one-bud one-leaf.
After comparing the content of organic acids in five different varieties of processed, roasted
green tea, it is found that the content of organic acids in Fuding species is the lowest
(0.538 mg/mL), the content of Huangjingui is the highest (0.716 mg/mL), and the content
of oxalic acid, quinic acid, and L-ascorbic acid in Yunkang 10 of large leaf species is higher
than those of medium and small leaf species. The content of organic acids in different kinds
of tea significantly changes, and the order is black tea > oolong tea > green tea > Pu ‘er
tea. Quinic acid, L-ascorbic acid, and citric acid are the main organic acids leading to the
difference. The processing technology, especially the degree of fermentation, is positively
correlated with the total content of organic acids. The lowest content of Pu ‘er tea may be
related to the tenderness of raw materials, post-fermentation process, and storage time. In
addition, the leaching of organic acids rises with the increase in brewing temperature and
brewing time but decreases with the increase in pH value [32,36].
Zhang et al. treated the Longjing 43 variety with a sunshade net and straws for
20 days and found that the content of organic acids in leaves and stems changed differently.
Compared with the control group, the relative contents of succinic acid and quinic acid in
leaves decreased, and malic acid, gallic acid, citric acid, and α-ketoglutarate increased. In
stems, quinic acid decreases under sunshade net cover and increases under straw cover [41].
Shirai found that the content of quinic acid was high in Kukicha and low in matcha. The
Molecules 2023, 28, 2339 7 of 11

quinic acid content increased with leaf maturity, as opposed to oxalic, malic, succinic, and
citric acids. Shading cultivation markedly reduced the quinic acid content and slightly
enhanced the content of malic, citric, and oxalic acids. Low-grade green tea is a good source
of quinic acid [42].

5. Organic Acids and Sensory Quality of Tea

Although organic acids only account for about 3% of the dry matter weight, it has
a great impact on the quality of tea leaves. Aromatic organic acids and aliphatic organic
acids in tea leaves contribute a lot to the formation of tea aroma and taste. Most of these
organic acids are soluble in water and are one of the main ingredients affecting the flavor
and quality of tea soup [4].

5.1. Organic Acids and Tea Aroma

The content of carboxylic acid in fresh tea leaves is lower than that in the finished tea.
One of the reasons for the flavor difference between black tea and green tea is the different
content and proportion of volatile organic acids. Studies have shown that the organic
acids account for about 30% of the total essential oils in black tea and only account for
2–3% in green tea. During the withering and fermentation process of black tea, unsaturated
fatty acids will be oxidized and degraded into volatile small molecules, such as aldehydes,
ketones, and acids, which then participate in the composition of black tea aroma [43].
During the processing of Keemun black tea, the content of most carboxylic acid aromatic
substances gradually increases, especially aromatic substances, such as cinnamic acid,
which constitutes the special aroma of Keemun black tea [35]. During the storage process
of tea leaves, the free fatty acids produced from the hydrolysis of lipid compounds are
the main reason for the musty and rancid tastes [44]. Under extreme environments, such
as high-temperature storage conditions, aerobic environments, and intense light, lipids
in tea leaves will be hydrolyzed or oxidized, resulting in the deterioration or aging of tea
leaves [45]. During the storage process of green tea, the content of organic acids such as
oleic acid and stearic acid, which are negatively correlated with the quality of tea aroma,
will increase. However, the content of organic acids, such as palmitic acid, lauric acid, and
linolenic acid, which are positively related to the quality of tea aroma, will decrease [46].

5.2. Organic Acids and Tea Flavor

The organic acids are one class of main components contributing to the sour taste of
tea soup, whose content is positively correlated with the degree of fermentation. Moderate
content of sour compounds increases the fullness of taste, but excessive content will lead
to the deterioration of quality [36,47]. Yue et al. found that sour taste was related to malic
acid, lactic acid, citric acid, and shikimic acid through the partial least squares and variable
projection analysis [48]. However, Zhang et al. reported that the acidity in black tea and
green tea was not directly related to organic acids but associated with the interaction
between organic acids and other substances, pH value, or other sour substances in tea
infusions [49].
The taste threshold is the critical concentration value at which each taste compound
can be perceived. The lower it is, the easier it is to be perceived. The taste activity value is
the ratio of the content to the taste threshold value. The higher the taste activity value is, the
greater the contribution of the compound to taste. Mao sums up the content range of taste
compounds in Gongfu black tea samples and calculates the taste activity value according
to the threshold value. In terms of sourness, L-malic acid, oxalic acid, and aspartic acid are
the main sour components of Gongfu black tea, followed by gallic acid and succinic acid.
This is different from the research results of Yue et al., which may be caused by different
tea varieties [38]. The taste characteristics and threshold values of common organic acids in
tea are shown in Table 3.
Molecules 2023, 28, 2339 8 of 11

Table 3. Taste characteristics and thresholds of common organic acids in tea.

Organic Acid Name Flavor Characteristics Threshold Value (mg·L−1 )

Gallic acid Sour, astringent 34.02
Oxalic acid Sour 45
Pyruvic acid Sour 130
L-malic acid Sour 87
L-ascorbic acid Sour 123.28
Lactic acid Sour 133.2
Acetic acid Sour 50
Citric acid Sour 770
Butanedioic acid Sour 106.28

Some scholars have proved that succinic acid can enhance the umami taste of amino
acid compounds [50,51]. Ascorbic acid, citric acid, succinic acid, and malic acid are natural
antioxidants that can lower the pH of tea soup, reduce the production of H2 O2 , and thus
maintain the flavor quality of tea soup [52]. Xu et al. researched the influence of Ca2+ on
the content of organic acids, turbidity, and sediment formation in green tea soup. The
results showed that the turbidity of tea soup is highly negatively correlated with the content
of oxalic acid, quinic acid, and tartaric acid [53]. Kaneko et al. are the first to find that
theogallin in matcha can enhance the umami taste, and so do succinic acid, and gallic
acid [54]. Through correlation analysis, Liu et al. [32] showed that the content of gallic acid
and succinic acid in high score group of green tea (characteristic of strong umami taste) is
also high and that the total amount of lactic acid, ascorbic acid, and organic acids has a
positive effect on the taste of tea soup [55]. Lv et al. analyzed the chemical components of
the taste quality of Pu ‘er tea and found that the content level of organic acids was markedly
negatively correlated with the score of taste quality [56]. Song et al. showed that fumaric
acid had a great contribution to the “mellow and fresh” taste style of black tea. All these
results indicate that organic acids play an important role in the taste of tea soup, but the
related mechanisms and influencing factors are complicated [57].

6. Health Benefits of Organic Acids

The organic acids in tea leaves can promote the human body’s absorption of catechin,
enhance the antioxidant properties of tea polyphenols, stimulate the activities of α-amylase
and trypsin, and promote the digestion and absorption of starch and protein [40,58–60]. In
addition, organic acids inhibit the growth of pathogenic intestinal bacteria and improve
intestinal function. In the pile-fermentation of Fuzhuan tea, a large number of organic
acids are produced, which can reduce the pH value in the intestine, inhibit the growth
and reproduction of pathogenic bacteria, and improve the gastrointestinal function of the
human body [61,62]. Dark tea extracts containing organic acids and other active ingredients
could accelerate gastrointestinal transit, promote colonization of probiotics, including
Bifidobacterium and Lactobacillus, and suppress the proliferation of harmful Enterococcus and
Escherichia coli in normal mice [63]. The antibacterial effect of organic acids in white tea
has been studied, and the results showed that all organic acids had inhibitory effects on
Staphylococcus aureus, Shigella flexneri, and enteric subspecies of Salmonella enterica, among
which mixed acid had the best antibacterial effect, followed by oxalic acid and acetic acid,
and fumaric acid had the worst effect [64].
Citric acid and malic acid are both important products of the tricarboxylic acid cycle.
Due to their special roles in metabolism, they can directly participate in metabolic activities
in the human body. Malic acid provides energy for the human body for a short period,
which can resist fatigue and protect the liver, kidneys, and heart, so it can be used to
develop healthy drinks. The addition of malic acid can increase drug stability and promote
the absorption and diffusion of the drug in the human body. In addition, L-malic acid has a
good antioxidant ability, which can delay the rancidity of food caused by oxidation and the
reduction of its nutritional value. Clinically, sodium citrate is used as an anticoagulant, and
Molecules 2023, 28, 2339 9 of 11

ferric ammonium citrate is a blood tonic. Citric acid is an important intermediate product of
the metabolism of carbohydrates, fat, and protein in animals and plants. In the aerobic state,
pyruvate is completely oxidized through the tricarboxylic acid cycle, which is the most
efficient way to obtain energy. In addition, salicylic acid, acetoacetic acid, palmitic acid, etc.,
also have antioxidant, heat-clearing, detoxification, sterilization, and other health benefits.

7. Summary and Prospect

Organic acids not only play an important role in the nutrient absorption of tea plants
but also contribute to the aroma, taste, and health benefits of tea. So far, it is known that
varieties, seasons, environmental conditions, such as light and soil pH, and processing
technology affect the composition and content of organic acids in tea plants and different
types of finished teas. Compared with other quality and functional ingredients in tea leaves,
there are fewer studies on the characteristics of organic acids in tea. This article reviewed the
research progresses of organic acids in tea, including their analysis methods, secretion and
physiological function of organic acids from tea tree rhizosphere, the composition of organic
acids in tea leaves and relevant influencing factors, the contribution of organic acids to
sensory quality, and their health benefits, such as antioxidation, promotion of digestion and
absorption, acceleration of gastrointestinal transit, and regulation of intestinal flora. The
in-depth study of organic acids is conducive to opening up new horizons in the selection
and breeding of tea varieties, the optimization of processing technology, and the research
on the health functions of tea.

Author Contributions: Writing—original draft preparation, F.Y., J.K., Q.L. and H.K.W.; writing—
review and editing, H.C. and B.L. All authors have read and agreed to the published version of
the manuscript.
Funding: This research was funded by the Talent Introduction Project of Zhejiang Shuren University
(KXJ0622604), the Natural Science Foundation of Zhejiang Province, China (LY21C200010), National
Undergraduate Innovation and Entrepreneurship Training Program (202211842014X), Shaoxing
Doctor Innovation Station (KHJ0622610), and School-Enterprise Cooperation Project (KHJ0622116).
Informed Consent Statement: Not applicable.
Conflicts of Interest: The authors declare no conflict of interest.

References
1. Du, J.; Bai, L.; Bai, B.Z. The major chemical constituents of tea. Agric. Technol. 2003, 23, 53–55.
2. Oburger, E.; Kirk, G.J.D.; Wenzel, W.W.; Puschenreiter, M.; Jones, D.L. Interactive effects of organic acids in the rhizosphere. Soil
Biol. Biochem. 2009, 41, 449–457. [CrossRef]
3. Wang, L.; Tang, J.; Xiao, B.; Yang, Y.; Yu, Y. Enhanced release of fluoride from rhizosphere soil of tea plants by organic acids and
reduced secretion of organic acids by fluoride supply. Acta Agric. Scand. Sect. B Soil Plant Sci. 2013, 63, 426–432. [CrossRef]
4. Tan, H.P.; Ye, S.R.; Chen, L.; Zou, Y. Determination overview of organic acids in tea. China Measuerment Test. Technol. 2008, 6,
77–80.
5. He, S.P.; Guo, C.F.; Sun, Y. Research progress of organic acids in tea. Subtrop. Agric. Res. 2015, 11, 63–67.
6. He, S.P.; Guo, C.F.; Sun, Y.; Zheng, D.Y. Determination of organic acids in White Tea by HPLC method. China Tea Process. 2015,
37–42. Available online: http://a.xueshu.baidu.com/usercenter/paper/show?paperid=ac1fdbc515484cb496d4475eb1732302
(accessed on 25 February 2023).
7. Lu, M.; Zhang, S.Y.; Yu, H.W.; Song, S.N. Analysis of organic acids in Maojian green tea by high performance liquid chromatogra-
phy. Guang Dong Agric. Sci. 2012, 39, 86–87.
8. Qu, Y.Q.; Marat, T.; Zheng, D.Y.; Zhang, Q.; Yang, J.F.; Ye, N.X. HPLC determination of phenolic acids in teas. Acta Tea Sin. 2019,
60, 69–74.
9. Wang, L.L.; Yang, J.G.; Lin, Q.X.; Xiang, L.H.; Song, Z.S.; Zhang, Y.G.; Chen, L. Determination of 10 organic acid contents in tea
using high performance liquid chromatography-diode array detector. J. Zhejiang Univ. (Agric. Life Sci.) 2019, 45, 47–53.
10. Zhang, J.; Jiang, H.J.; Liu, Z.H.; Chen, J.H.; Lin, Y.; Zou, W.M. Simultaneous separation and determination of organic acids in
black tea by high performance liquid chromatography using polar-enhance C18 column. J. Food Saf. Qual. 2014, 5, 2476–2481.
11. Moldoveanu, S.C.; Poole, T.; Scott, W.A. An LC-MS Method for the Analysis of Some Organic Acids in Tobacco Leaf, Snus, and
Wet Snuff. Contrib. Tob. Nicotine Res. 2018, 28, 30–41. [CrossRef]
Molecules 2023, 28, 2339 10 of 11

12. Song, W.-S.; Park, H.-G.; Kim, S.-M.; Jo, S.-H.; Kim, B.-G.; Theberge, A.B.; Kim, Y.-G. Chemical derivatization-based LC–MS/MS
method for quantitation of gut microbial short-chain fatty acids. J. Ind. Eng. Chem. 2020, 83, 297–302. [CrossRef]
13. Awad, H.; Allen, K.J.H.; Adamko, D.J.; El-Aneed, A. Development of a new quantification method for organic acids in urine as
potential biomarkers for respiratory illness. J. Chromatogr. B 2019, 1122–1123, 29–38. [CrossRef] [PubMed]
14. Xu, J.; Hu, F.-L.; Wang, W.; Wan, X.-C.; Bao, G.-H. Investigation on biochemical compositional changes during the microbial
fermentation process of Fu brick tea by LC–MS based metabolomics. Food Chem. 2015, 186, 176–184. [CrossRef]
15. Sandín-España, P.; Mateo-Miranda, M.; López-Goti, C.; De Cal, A.; Alonso-Prados, J.L. Development of a rapid and direct method
for the determination of organic acids in peach fruit using LC–ESI-MS. Food Chem. 2016, 192, 268–273. [CrossRef] [PubMed]
16. Fraser, K.; Lane, G.A.; Otter, D.E.; Harrison, S.J.; Quek, S.-Y.; Hemar, Y.; Rasmussen, S. Non-targeted analysis by LC–MS of major
metabolite changes during the oolong tea manufacturing in New Zealand. Food Chem. 2014, 151, 394–403. [CrossRef]
17. Han, W.Q.; Wang, B.; Wang, L.; Xian, Y.P. Quantitative analysis of organic acid and flavonoid in Herbal Tea by UPLC-MS/MS.
Food Res. Dev. 2017, 38, 130–134.
18. Lu, C.G.; Yang, J.; Kang, C.Z.; Yang, W.Z.; Zhou, L.; Li, Z.H.; Guo, L.P. Determination of nine organic acids in dendrobium
officinale by UPLC-MS/MS. J. Chin. Med. Mater. 2017, 40, 1360–1363.
19. Wang, S.W.; Liu, Y.P.; Wang, X.N.; Sun, H.B. Determination of ten organic acids content in fresh litchi based on high performance
liquid chromatography with triple quadrupole mass spectrometry technique. Chin. J. Pestic. Sci. 2019, 21, 359–365.
20. Morita, A.; Yanagisawa, O.; Maeda, S.; Takatsu, S.; Ikka, T. Tea plant (Camellia sinensis L.) roots secrete oxalic acid and caffeine into
medium containing aluminum. Soil Sci. Plant Nutr. 2011, 57, 796–802. [CrossRef]
21. Wang, X.P. Study on the organic acids excreted by tea roots. J. Tea Sci. 1994, 14, 17–22.
22. Pang, X.; Wang, Y.H.; Wang, W.D.; Yin, Y.; Shu, Z.; Chen, X.; Li, X.H. Effects of tea pruning materials and tea polyphenols on
organic acids secretion and mineral uptake in tea plant. J. Tea Sci. 2014, 34, 591–600.
23. Tang, D.D.; Liu, M.Y.; Zhang, Q.F.; Shi, Y.Z.; Ma, L.F.; Ruan, J.Y. Effects of nitrogen form and root-zone pH on nutrient uptake and
concentrations of organic anions in tea plants (Camellia sinensis). J. Tea Sci. 2019, 39, 159–170.
24. Ruan, J.Y.; Yang, Y.J.; Ma, L.F. Recent progress in the research on fluoride in tea:accumulation characteristics by plants and
concentrations in products with regarding to the impact on human health. J. Tea Sci. 2007, 27, 1–7.
25. Luo, X.P.; Li, L.X.; He, C.L. Advanced study on fluoride and measures for reduction of fluoride content in tea (Camellia sinensis).
Tea Sci. Technol. 2006, 2, 6–9.
26. Liu, T.T.; Zhao, Q.; Gao, H.J.; Wan, X.C.; Zhang, Z.Z. Effects of organic acids and iron plaque outside roots on absorption and
accumulation of fluoride in tea plants. J. Nanjing Agric. Univ. 2013, 36, 72–78.
27. Liu, T.T.; Gao, H.J.; Wan, X.C.; Zhang, Z.Z. Impacts of aluminum on root cell membrane permeability and organic acids in root
exudates of tea plant. J. Tea Sci. 2011, 31, 458–462.
28. Morita, A.; Fujii, Y.; Yokota, H. Effect of aluminium on exudation of organic acid anions in tea plants. In Plant Nutrition:
Food Security and Sustainability of Agro-Ecosystems through Basic and Applied Research; Horst, W.J., Schenk, M.K., Bürkert, A.,
Claassen, N., Flessa, H., Frommer, W.B., Goldbach, H., Olfs, H.W., Römheld, V., Sattelmacher, B., et al., Eds.; Springer: Amsterdam,
The Netherlands, 2001; pp. 508–509.
29. Lin, H.T.; Shi, Y.X. Effects of Pb and Cd stress on organic acids secreted by tea roots. Shandong Agric. Sci. 2005, 2, 32–34.
30. Zeng, Q.R.; Zhou, X.H.; Liao, B.H.; Yang, R.B.; Tie, B.Q. Activation effects of low-molecular-weight organic acids on Al, F, P, Cu,
Zn, Fe and Mn in soils of tea garden. J. Tea Sci. 2001, 21, 48–52.
31. Marat, T.; Zhang, Q.; Qu, Y.Q.; Zheng, D.Y.; Yang, J.F.; Ye, N.X. Analysis of organic acid components in white tea processing. Tea
Fujian 2019, 41, 11–12.
32. Liu, P.P.; Zheng, P.C.; Guo, G.Y.; Jin, X.F.; Yin, P.; Gao, S.W.; Wang, S.P.; Ye, F.; Zheng, L.; Gong, Z.M. Effects of tea varieties and
growing period on organic acid content in tea leaves. Jiangsu Agric. Sci. 2018, 46, 225–228.
33. Xie, J.M.; Luo, M.Y.; Liu, Y.F.; Zhu, Q. Quality analysis for aged Keemun Black Tea. Hunan Agric. Sci. 2012, 21, 100–102, 105.
34. Yuan, L.; Huang, J.A.; Gong, Z.H.; Chen, J.H.; Li, Y.H.; Yuan, Y.; Xiao, W.J. Study on development and application technology of
analysis method for organic acid in Fu-brick Tea by RP-HPLC. Chin. Agric. Sci. Bull. 2011, 27, 246–252.
35. Zhao, H.T. Metabolism of organic acids in black tea processing and its effect on tea aroma formation. J. Tea Commun. 1993, 1,
25–27.
36. Liu, P.P.; Zhong, X.Y.; Xu, Y.Q.; Chen, G.S.; Yin, J.F.; Liu, P. Study on organic acids contents in tea leaves and its extracting
characteristics. J. Tea Sci. 2013, 33, 405–410.
37. Yu, F.; Chen, C.; Chen, S.; Wang, K.; Huang, H.; Wu, Y.; He, P.; Tu, Y.; Li, B. Dynamic changes and mechanisms of organic acids
during black tea manufacturing process. Food Control 2022, 132, 108535. [CrossRef]
38. Mao, S.H. Quality Analysis and Control of Congou Black Tea Based on Sensomics. Ph.D. Thesis, Southwest University, George-
town, TX, USA, 2018.
39. Li, S.; Gong, X.; Liu, Z.H.; Huang, H.; Huang, J.A. The qualitative component conversion in dark tea after colonization of Eurotium
cristatum strains. Mycosystema 2014, 33, 713–718.
40. Ding, L. Studies on the Changes of Main Chemical Components and it’s Activation to the Pancreatin in the Processing of Fuzhuan
Brick Tea. Master’s Thesis, Zhejiang University, Hangzhou, China, 2006.
41. Zhang, Q.; Liu, M.; Mumm, R.; Vos, R.C.H.; Ruan, J. Metabolomics reveals the within-plant spatial effects of shading on tea plants.
Tree Physiol. 2021, 41, 317–330. [CrossRef]
Molecules 2023, 28, 2339 11 of 11

42. Shirai, N. Organic acid analysis in green tea leaves using high-performance liquid chromatography. J. Oleo Sci. 2022, 71, 1413–1419.
[CrossRef]
43. Liao, S.J.; Tong, H.R. Studies on the fatty acid and glycoside aroma precursors of different tea varieties. J. Southwest Univ. 2008, 30,
62–66.
44. Zhang, Z.Z.; Shi, Z.P.; Wan, X.C. Changes of fatty acids during storage of green tea and their effects on aroma. China Tea Process.
1999, 2, 39–41.
45. Cui, F. Study on the Change of Main Quality Components and Its Influence Factors during Green Tea Storage. Master’s Thesis,
Zhejiang University, Hangzhou, China, 2008.
46. Wu, X.C. Free fatty acid composition in green tea storage. J. Tea Commun. 1990, 23–24.
47. Wang, M.L. Discusses the reason and the control method of the acidness of the black tea. Tea Fujian 2014, 36, 32–33.
48. Yue, C.N.; Qin, D.D.; Cai, H.L.; Wang, Z.H.; Li, C.; Li, Y.S.; Yang, P.X. Taste characteristics and key compounds analysis of Congou
black tea in northern Jiangxi Province. Food Ferment. Ind. 2021, 47, 260–267.
49. Zhang, X.; Du, X.; Li, Y.Z.; Nie, C.N.; Wang, C.M.; Bian, J.J.; Luo, F. Are organic acids really related to the sour taste difference
between Chinese black tea and green tea? Food Sci. Nutr. 2022, 10, 2071–2081. [CrossRef] [PubMed]
50. Wu, N.; Gu, S.Q.; Tao, N.P.; Wang, X.C. Research progress in interaction between umami substances. Food Ind. Technol. 2014, 35,
389–392, 400.
51. Zhang, L.; Cao, Q.-Q.; Granato, D.; Xu, Y.-Q.; Ho, C.-T. Association between chemistry and taste of tea: A review. Trends Food Sci.
Technol. 2020, 101, 139–149. [CrossRef]
52. Aoshima, H.; Ayabe, S. Prevention of the deterioration of polyphenol-rich beverages. Food Chem. 2007, 100, 350–355. [CrossRef]
53. Xu, Y.-Q.; Zhong, X.-Y.; Yin, J.-F.; Yuan, H.-B.; Tang, P.; Du, Q.-Z. The impact of Ca2+ combination with organic acids on green tea
infusions. Food Chem. 2013, 139, 944–948. [CrossRef]
54. Kaneko, S.; Kumazawa, K.; Masuda, H.; Henze, A.; Hofmann, T.F. Molecular and sensory studies on the umami taste of Japanese
green tea. J. Agric. Food Chem. 2006, 54, 2688–2694. [CrossRef]
55. Liu, S.; Yang, T.; Tan, J.F.; Lu, H.P.; Guo, L.; Lin, Z.; Lu, C.Y. Quantitative descriptive analysis and its correlation between chemical
compoments of taste in green tea. Chin. Agric. Sci. Bull. 2014, 30, 40–46.
56. Lu, H.P.; Zhang, Y.; Yang, T.; Shi, J.; Lin, Z. Analysis on chemical constituents of taste quality of Pu ‘er tea. Food Ferment. Ind. 2016,
42, 178–183.
57. Song, C.J.; Fan, F.Y.; Gong, S.Y.; Guo, H.W.; Li, C.L.; Zong, B.Z. Taste characteristic and main contributing compounds of different
origin black tea. Sci. Agric. Sin. 2020, 53, 383–394.
58. Dufresne, C.; Farnworth, E. Tea, Kombucha, and health: A review. Food Res. Int. 2000, 33, 409–421. [CrossRef]
59. Wang, Y. The anti-oxidation and anti-microbial activities of tea polyphenols and its increased reagents. J. Biol. 2007, 24, 54–56.
60. Chen, W.F.; Tu, Y.Y.; Wu, Y.Y.; Liang, H.L.; Wang, Y. Effect of compacted tea extract of dark tea on trypsin activity. China Tea 2002,
3, 16–17.
61. Wu, X.L. Experimental Study of Dark Tea Improving Gastrointestional Function in Mice. Master’s Thesis, Hunan Agricultural
University, Changsha, China, 2013.
62. Tu, Y.Y.; Xu, H.R.; Liang, H.L.; Chen, X.; Wu, Y.Y.; Chen, W.F.; Tan, Q.F. Effects of compacted tea on trypsin activity and beneficial
bacteria in intestinal tract. Food Sci. 2002, 10, 113–116.
63. Gong, Z.P.; Ouyang, J.; Wu, X.L.; Zhou, F.; Lu, D.M.; Zhao, C.J.; Liu, C.F.; Zhu, W.; Zhang, J.C.; Li, N.X.; et al. Dark tea extracts:
Chemical constituents and modulatory effect on gastrointestinal function. Biomed. Pharmacother. 2020, 130, 110514. [CrossRef]
64. He, S.P. Study on Organic Acids of White Tea and Their Antibacterial Effects In Vitro. Master’s Thesis, Fujian Agriculture and
Forestry University, Fuzhou, China, 2016.

Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual
author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to
people or property resulting from any ideas, methods, instructions or products referred to in the content.